A New Species of the Armored Catfish Genus Pareiorhaphis Miranda Ribeiro (Siluriformes: Loricariidae) from the Rio Paraguaçu, Bahia State, Northeastern Brazil

A new species of the armored catfish genus Pareiorhaphis Miranda Ribeiro (Siluriformes: Loricariidae) from the rio Paraguaçu, Bahia State, northeastern Brazil

Edson H. L. Pereira1 and Angela M. Zanata2

A new armored catfish species of the genus Pareiorhaphis is described from the middle and upper portions of rio Paraguaçu basin, coastal drainage of Bahia State, northeastern Brazil. The new species is readily distinguished from all its congeners by having two putative autapomorphies: (1) skin fold just posterior to each emergent tooth series of dentary formed by a single enlarged, flattened papilla, and (2) the midline of lower lip immediately behind the dentaries with small patch of distinct papillae arranged in a short median bump. In addition, the shallow caudal peduncle and comparatively lower number of teeth in each dentary also distinguishes the new species from all congeners. The new species is also compared to Pareiorhaphis bahianus, the geographically closest congener. Uma espécie nova de cascudo do gênero Pareiorhaphis é descrita da porção média e superior da bacia do rio Paraguaçu, drenagem costeira do estado da Bahia, nordeste do Brasil. A espécie nova é facilmente diagnosticada das demais congêneres por apresentar duas possíveis autapomorfias: (1) uma prega de pele atrás de cada série emergente de dentes do dentário formada por uma única papila larga e achatada e (2) um conjunto de papilas distintas arranjadas em uma elevação curta localizada na linha média do lábio inferior. Além disso, a menor altura do pedúnculo caudal e o baixo número de dentes em cada dentário também distinguem a espécie nova de todas as congêneres. A espécie nova é ainda comparada com Pareiorhaphis bahianus, congênere geograficamente mais próximo. Key words: Cascudos, Chapada Diamantina, Neoplecostominae, Neoplecostomus, Pareiorhaphis bahianus.

Introduction to southern drainages, with at least nine species occurring in Santa Catarina State, in the last two decades six new species Pareiorhaphis Miranda Ribeiro currently comprises were revealed to occur in rivers draining a more northeastern 21 nominal valid species of small to medium-sized portion of the country (e.g., rio Doce, rio Jequitinhonha, and suckermouth armored catfishes easily recognized by a rio São Francisco drainages), and two small coastal rivers of series of derived features, including cheeks, opercle, and the Espírito Santo State, named Piraquê-Açu and Reis Magos. the exposed lateral process of the cleithrum of adult males Up to date, P. bahianus (Gosline) described from Fazenda covered with hypertrophied odontodes, and the lateroventral Almada, Ilhéus, is the only species known to occur in Bahia portion of preopercle deeply rugose due to the attachment State and represents the northernmost distribution record of of hypertrophied odontodes (Pereira et al., 2007, 2010). the genus. The type locality of P. bahianus is within the area Species of Pareiorhaphis are almost completely restricted drained by the rio Almada and this species is recorded from to rivers draining the Brazilian Shield, with the exception various small coastal rivers located between the rio Pardo and of P. regani (Giltay), which was described from a tributary rio de Contas basins (Carvalho & Bockmann, 2007). of the rio Negro, in the Amazon basin. Although the highest Field expeditions over the last eight years in coastal species diversity of the genus has been traditionally attributed drainages of Bahia State yielded several undescribed species

1Laboratório de Sistemática de Vertebrados, Pontifícia Universidade Católica do Rio Grande do Sul. Av. Ipiranga, 6681, P. O. Box 1429, 90619-900 Porto Alegre, RS, Brazil. [email protected] 2Departamento de Zoologia, Instituto de Biologia, Universidade Federal da Bahia. Rua Barão de Geremoabo, s/n, Ondina, 40170-290 Salvador, BA, Brazil. [email protected] 35 36 New Pareiorhaphis from northeastern Brazil of armored catfishes, revealing an unknown diversity of Paratypes. All from Brazil, Bahia State, rio Paraguaçu basin. MCP loricariids in the area. This fact can be exemplified by 47712, 18, 30.1-82.1 mm SL (6, 63.1-82.1 mm SL), 1 c&s, 64.1 mm SL, and UFBA 7063, 25, 31.0-71.8 mm SL (3, 66.5-71.8 mm SL), discoveries of two new species of Hypostomus describe collected with the holotype. MCP 48004, 3, 72.6-81.8 mm SL, 1 c&s, recently (Birindelli et al., 2007; Zanata et al., 2013) and a 72.6 mm SL, rio Paraguaçu at Fazenda Os Touros, ca. 18 km from new species of Pareiorhaphis described herein. Iaçu, 12°41’10”S 40°7’47”W, 143 m a.s.l., Sep 2011, J. J. S. Conceição. MZUSP 86089, 3, 45.5-57.8 mm SL, rio Paraguaçu, upstream Itaetê, Material and Methods 12°58’25”S 40°58’20”W, 21 Sep 2004, M. E. Bichuette et al. MZUSP 86154, 1, 62.8 mm SL, rio Paraguaçu, upstream Itaetê, 12°58’25”S 40°58’20”W, 8 Nov 2004, M. E. Bichuette & F. D. Passos. MZUSP All morphometric features were taken as point- 88163, 10, 54.8-71.4 mm SL; UFBA 7026, 5, 54.2-62.5 mm SL, rio to-point linear distances with digital calipers under a Paraguaçu at Fazenda Os Touros, ca.18 km from Iaçu, 12°41’10”S stereomicroscope, and recorded to the nearest 0.1 mm. 40°7’47”W, 143 m a.s.l., 8 Jun 2005, A. M. Zanata, J. L. O. Birindelli, Counts and measurements followed Pereira et al. (2007). O. T. Oyakawa, M. P. Geraldes, P. Camelier & P. Moura. UFBA 6188, 1, 79.3 mm SL, rio Paraguaçu at Fazenda Os Touros, ca. 18 km from Nomenclature and counts of body plates follow Schaefer Iaçu, 12°41’10”S 40°7’47”W, 15 May 2010, A. M. Zanata, R. Burger, (1997). Standard length (SL) is expressed in millimeters C. Primitivo, D. França & J. J. S. Conceição. UFBA 7350, 6, 30.5- while all other measurements are given as percents of SL, 80.9, rio Una at Cachoeira das Andorinhas, Iramaia, 13°19’52.5”S except for subunits of the head, which are presented as 41°6’53.4”W, 449 m a.s.l., 1 May 2013, A. M. Zanata, R. Burger, L. percents of the head length (HL). Counts of procurrent Oliveira & R. Abreu. caudal-fin rays and vertebrae were made only in cleared and counterstained specimens (c&s) prepared according Diagnosis. Pareiorhaphis lophia differs from all congeners to Taylor & Van Dyke (1985). Vertebral counts comprise in having two exclusive features: (1) a skin fold just posterior all vertebrae including the five centra modified into the to each emergent tooth series of dentary formed by a single Weberian apparatus and the compound caudal centrum enlarged, flattened papilla, and (2) the midline of lower lip (PU1+U1) was counted as one element. Meristic and immediately behind the dentaries with small patch of distinct morphometric data for comparisons of Pareiorhaphis species papillae arranged in a short median bump (vs. papillae not in the diagnosis section are from Pereira & Reis (2002), projected in a bump). Moreover, the lower caudal peduncle Pereira (2005), Pereira et al. (2007, 2010), Pereira & Britto also distinguishes P. lophia from all remaining Pareiorhaphis (2012), and Pereira et al. (2012). Institutional abbreviations species (6.7-7.8 vs. 7.8-14.0% SL), with the exception of are: Museu de Ciências e Tecnologia, Pontifícia Universidade P. proskynita. Furthermore, P. lophia can be distinguished Católica do Rio Grande do Sul, Porto Alegre (MCP); from remaining congeners but P. bahianus, P. cerosus, P. Museu de Zoologia, Universidade de São Paulo, São Paulo eurycephalus, P. nudulus, P. parmula, P. steindachneri, and (MZUSP); and Universidade Federal da Bahia, Salvador P. vestigipinnis by the lower number of dentary teeth (20- (UFBA). Descriptions of coloration were based on specimens 38 vs. 42-120). Pareiorhaphis lophia can be distinguished preserved in ethanol and photographs of the freshly collected from P. bahianus, P. cerosus, P. eurycephalus, P. parmula, specimens. In the lists of examined material the museum and P. steindachneri by the usual absence of first dorsal-fin abbreviation and catalog number come first, followed by the spinelet (vs. spinelet present). The new species can be further number and SL range of specimens in that lot, the number distinguished from P. nudulus and P. vestigipinnis by the and SL range of specimens measured for the morphometric possession of an adipose fin (vs. adipose fin absent) and the comparisons, in parentheses, and locality. narrower caudal peduncle (4.1-5.3 vs. 5.5-8.7% SL). Adult males of Pareiorhaphis are herein defined according to Pereira et al. (2007). The specimens included in the list of Description. Counts and measurements in Table 1. Small to material examined are a combination of male, female and medium-sized loricariid with standard length of measured immature specimens of both sexes. specimens 45.5-82.1 mm SL. Head and body moderately depressed. Greatest body width at cleithrum, progressively Results tapering to end of caudal peduncle. Dorsal profile of body slightly convex, from snout tip to dorsal-fin origin and Pareiorhaphis lophia, new species almost straight from that point to end of adipose fin, then Figs. 1-3 slightly concave to caudal fin. Greatest body depth at dorsal- fin origin. Least body depth at shallowest part of caudal Holotype. MCP 47711, 76.9 mm SL, male, Brazil, Bahia State, peduncle. Trunk and caudal peduncle mostly oval in cross- Lençóis, rio Paraguaçu drainage, rio Santo Antônio at bridge on road section, flattened ventrally and more compressed caudally. BR-242, 12°29’37”S 41°19’38”W, 350 m above sea level, 7 Jul 2011, Lateral-line canal in median series complete, pored tube A. M. Zanata, P. Camelier, J. Birindelli, R. Burger & B. Sardeiro. visible from compound pterotic to caudal-fin base. Ventral E.H.L. Pereira & A.M. Zanata 37

Fig. 1. Pareiorhaphis lophia, MCP 47711, holotype,76.9 mm SL, male, Brazil, Bahia State, Lençóis, rio Paraguaçu drainage, rio Santo Antônio at bridge on road BR-242, 12°29’37”S 41°19’38”W. profile almost straight between snout tip and pelvic girdle, series of pairs in some specimens, or without any noticeable slightly elevating posteriorly along anal-fin base, almost arrangement. Five lateral rows of dermal plates covering straight along caudal peduncle. Dorsal surface of body body, not forming keels; mid-dorsal and mid-ventral series covered by plates except for narrow naked area around of lateral plates incomplete, terminating four or five plates dorsal-fin base. Predorsal plates arranged in three to four before caudal fin. Abdomen entirely devoid of plates. First 38 New Pareiorhaphis from northeastern Brazil anal-fin pterygiophore covered by skin. Lower surface of Dorsal fin origin on vertical line through pelvic-fin head and portion from pelvic-fin insertions to anal-fin origin origin. Dorsal fin short, reaching three plates behind its base completely naked, except by plated region around anal fin. terminus when adpressed. Nuchal plate exposed, not covered Head broad and moderately depressed. Outline of head by skin. Dorsal-fin spinelet usually absent. Dorsal-fin locking round in dorsal view. Interorbital space straight or slightly mechanism non-functional. Dorsal-fin spine moderately concave. Three weakly elevated ridges between orbits and flexible, followed by seven branched rays. Distal margin of snout tip. Median ridge slightly more prominent. Snout dorsal fin straight. Adipose fin with well-ossified leading spine convex in lateral profile; snout tip with rounded area of bearing odontodes. Adipose-fin membrane short or extended naked skin, devoid of odontodes. Adult males with well- slightly beyond adipose-fin spine. Adipose fin preceded by two developed soft fleshy lobes on lateral portion of head. or three median azygous plates. Pectoral-fin origin situated Soft fleshy area ornamented with short hypertrophied more dorsally than pelvic-fin origin. Pectoral fin moderate in odontodes, approximately perpendicular to body axis (Fig. size, with curved and flattened unbranched ray, covered by 1). Eye small, dorsolaterally placed. Iris operculum small minute odontodes in immature males and females. Adult males or absent in some specimens. Nares ovoid, slightly longer with first pectoral-fin spine very broad and straight, curved on than wide, positioned much closer to anterior margin of its base; bearing straight and short hypertrophied odontodes on orbit than to snout tip. its entire surface; dorsal surface with discrete dermal flap along Oral disk circular. Lips moderately developed, occupying its entire length. Pectoral fin I,6; first and second branched most of ventral surface of head. Upper lip narrow, bearing rays distinctly longer than spine and subsequent branched rays small rounded papillae. Lower lip wide but not reaching decreasing gradually in size. Posterior margin of pectoral fin pectoral girdle. Lower lip densely covered by minute straight, surpassing pelvic-fin origin when adpressed. Pelvic papillae. Papillae surrounded by small naked areas, fin i,5, not reaching or just reaching origin of anal fin when decreasing in size towards edge. Patch of distinct papillae at adpressed. Pelvic-fin unbranched ray depressed, covered with midline of lower lip immediately behind dentaries, arranged minute odontodes ventrally and laterally. Well-developed in short median and forming a bump (Fig. 2). Posterior dermal flap on dorsal surface of pelvic-fin unbranched ray; flap margin of lower lip fringed. Maxillary barbel short, mostly of males distinctly higher near fin base, and extending to tip adnate to lower lip and with small free portion distally. Teeth of ray in adult males; flap of females reduced or absent. Anal series in both premaxillae and dentaries forming shallow arc fin i,5. Caudal-fin posterior margin forked or slightly concave, with overall angle less than 180 degrees, with mesial ends asymmetrical; lower lobe longer than upper; 14 branched rays. slightly curved inwards. Teeth slender, asymmetrically bifid, Upper caudal-fin lobe with four and lower lobe with two or medial cusp slightly curved inwards. Lateral cusp long and three plate-like procurrent rays. Posteriormost procurrent rays pointed, reaching half-length of medial cusp. Distinct skin elongate. Hypural plate asymmetrical with lower lobe longer fold posterior to each emergent tooth series of dentary formed than upper. Total vertebral centra 28. by single enlarged, flattened papilla (Fig. 2). Color in alcohol. Ground color of dorsal surface of head and body light to dark brown; yellowish or whitish pale ventrally (Fig. 1). Cheeks, anterior margin of snout, area between nostrils and orbits, and compound pterotic covered with small, irregularly shaped, light or dark brown blotches. Soft fleshy skin on lateral portion of head well developed and whitish in adult males. Narrow transversal light brown bar extending through middle portion of supraoccipital. Dorsum and most of flank covered by irregularly closely- set darker brown blotches sometimes coalescing. Blotches usually arranged in five inconspicuous transverse saddles, situated between posterior margin of supraoccipital and origin of dorsal fin, behind dorsal-fin spine, posterior portion of dorsal-fin base, at preadipose area, and between adipose and caudal fins. Midlateral region of body with dark brown blotches contrasting with somewhat reticulate light areas. Fig. 2. Lower lip detail of Pareiorhaphis lophia, paratype, MCP Ventral portion of head and abdomen either completely 47712, 71.1 mm SL. Scale bar = 1 mm. Arrow on top indicates whitish or pale yellow. Ventral portion of caudal peduncle the flattened papilla and arrow on bottom indicates the bump. with numerous small dark blotches more conspicuous in E.H.L. Pereira & A.M. Zanata 39

Table 1. Morphometric and meristic data of the type series of Pareiorhaphis lophia. Values given as percents of standard length or head length. H = holotype, n = number of specimens, and SD = standard deviation. H n Low High Mean SD Standard length (mm) 76.9 33 45.5 82.1 65.1 - Percents of standard length Head length 34.9 33 31.4 36.2 34.0 1.25 Predorsal length 43.5 33 40.0 44.3 42.7 1.05 Postdorsal length 40.5 33 36.5 43.7 41.0 1.52 Preanal length 63.9 33 61.0 68.0 64.3 1.92 Preadipose length 80.1 33 77.6 82.4 80.2 1.10 Dorsal-fin spine length 21.0 33 19.3 22.8 21.1 0.80 Anal-fin unbranched ray length 14.1 33 11.9 15.0 13.4 0.77 Pectoral-fin spine length 17.4 33 16.4 21.7 18.9 1.27 Ventral-fin unbranched ray length 24.5 33 21.5 25.7 23.7 1.15 Upper principal caudal-fin ray 21.3 30 19.4 25.5 22.9 1.75 Lower principal caudal-fin ray 26.0 30 22.6 30.6 26.9 1.99 Adipose-fin spine length 8.9 33 7.1 10.2 9.0 0.73 Adipose to caudal fin distance 20.2 33 17.9 21.4 19.4 0.86 Trunk length 16.9 33 15.9 18.9 17.6 0.74 Abdominal length 22.7 33 21.3 24.8 22.9 0.85 Cleithral width 29.4 33 27.4 30.7 28.8 0.85 Body depth at dorsal-fin origin 17.4 33 14.0 18.9 16.6 1.37 Body width at dorsal-fin origin 21.2 33 19.1 24.7 22.1 1.42 Body width at anal-fin origin 14.1 31 12.2 15.4 14.0 0.82 Caudal peduncle length 34.9 33 33.4 39.4 36.2 1.56 Caudal peduncle depth 7.7 33 6.7 7.8 7.2 0.33 Caudal peduncle width 5.1 33 4.1 5.3 4.7 0.30 Percents of head length Snout length 62.4 33 58.1 65.1 61.9 1.47 Orbital diameter 13.2 33 11.8 15.7 13.9 0.94 Interorbital width 26.3 33 25.2 29.8 27.7 1.27 Head depth 48.9 33 42.4 52.8 47.1 2.81 Mandibular ramus left 18.7 33 15.1 19.7 17.1 1.30 Mandibular ramus right 19.2 33 15.1 19.3 17.0 1.33 Counts Premaxillary teeth left 40 30 23 40 30.8 4.39 Premaxillary teeth right 37 31 21 40 30.9 5.04 Dentary teeth left 32 30 20 38 26.9 4.85 Dentary teeth right 28 30 21 38 27.3 4.61 Plates in median lateral series left 27 33 23 27 25.0 1.02 Plates in median lateral series right 28 32 24 28 25.3 1.08 Plates at dorsal-fin base 6 33 5 7 6.0 0.50 Plates between dorsal and adipose 6 33 6 8 6.5 0.56 Plates between adipose and caudal 4 33 3 5 3.4 0.56 Plates at anal-fin base 2 33 2 3 2.5 0.51 Plates between anal and caudal fins 12 33 10 12 11.2 0.50 Preadipose azygous plates 2 33 2 3 2.3 0.48 region after posterior margin of anal fin. First thickened rays Distribution. The new species is known from middle portion of dorsal, pectoral, and pelvic fins light brown or grayish. of the rio Paraguaçu and two tributaries of its upper portion Dorsal and caudal fin branched rays with 2-5 transverse (Fig. 4a). Extensive ichthyological collecting efforts through irregular dark bars. Paired fins with diffuse blotches from the main coastal rivers across Bahia in recent years failed to middle to margin of fin, sometimes arranged in few irregular yield samples of Pareiorhaphis lophia from outside the rio series. One or two dark brown blotches at adipose-fin spine. Paraguaçu drainage, making it possible that the species is endemic to that river basin. Color in life. Color pattern of freshly collected specimens light brown to yellowish tan with small and irregular dark Common name. Acari, chupa-pedra. spots scattered on dorsal region of head and predorsal area (Fig. 3). Posterior region of body and flank with wide dark Habitat and ecological notes. The rio Paraguaçu is a brown transversal saddles. Soft fleshy skin on lateral portion blackwater coastal river situated in northeastern Brazil, with of head and first thickened rays of pectoral and pelvic fins headwaters on highlands of central Bahia State. The river yellowish. Markings on caudal fin conspicuous with narrow stretch in Iaçu, at the period of sampling, was around 60 m blotches irregularly arranged on unbranched and branched wide with rocky platforms exposed on the river borders. The caudal-fin rays sometimes forming four or five dark brown riparian vegetation is represented by a few trees restricted to narrow stripes. Iris golden. the river border and the surrounding area is almost completely 40 New Pareiorhaphis from northeastern Brazil

Fig. 3. Lateral view of Pareiorhaphis lophia, holotype, MCP 47711, 76.9 mm SL, photographed immediately after collection (left side of body). Photo by J. L. O. Birindelli. anthropized mainly by cattle ranching. The specimens that is present in females but reduced. In adult males, the were collected at night, on the shallow rocky platforms thickened first pelvic-fin unbranched ray have developed and inside crevices. Data about the other location on the dermal flap along the entire dorsal surface and distinctly rio Paraguaçu, situated upstream Itaetê, are not available. higher near fin base, while in females the dermal flap is The rio Santo Antônio, a blackwater tributary of the upper reduced. Compared to males, females possess an enlarged, rio Paraguaçu, is around 20 meters wide and surrounded by swollen urogenital opening, while males have a small and riparian forest in the stretch sampled (Fig. 4b). Stretches pointed urogenital papilla. of rapids and pools alternate, with rocks, loose stones, pebbles, and sand composing the substrate. At that sampling Comments. The absence of first dorsal-fin spinelet also point, the specimens of Pareiorhaphis lophia were hidden diagnoses Pareiorhaphis lophia from most of the remaining among pebbles on the shallow rapids. In the rio Una, also congeners. However, variation in this feature was noted in the a blackwater river, the new species was sampled on narrow type series. The single specimen of the lot MCP 47712 (64.1 stretches around 50 cm deep, just below a waterfall, with fast mm SL) has a reduced first dorsal-fin spinelet. Although the water current and rocky bottom. The specimens gathered at absence of dorsal fin-spinelet must be considered diagnostic this locality were hidden in crevices or under large stones. to Pareiorhaphis lophia, the variation related to this feature The new species occurs syntopically with the loricariids deserves mention. Hypostomus chrysostiktos Birindelli, Zanata & Lima and H. jaguar Zanata, Sardeiro & Zawadzki. Etymology. The species name lophia is from the Greek Λοϕια, meaning crest, ridge. In allusion to the presence of Sexual dimorphism. The sexual dimorphism of distinct bump on the lower lip, a diagnostic feature of the Pareiorhaphis lophia is characterized mainly by the large new species. A noun in apposition. soft fleshy lobes restricted to middle portion of the lateral margins of head that extends from anterior margin of nostrils Discussion to posterior margin of orbits of adult males. This soft area is ornamented by short hypertrophied odontodes. Odontodes The most distinctive feature of Pareiorhaphis lophia is also occur in females and juveniles, but are comparatively the single narrow and transversally flattened papilla, just much smaller, while soft lobes are absent in females and posterior to each dentary, which easily distinguishes P. juveniles. Also, adult males have the unbranched pectoral lophia from all its congeners. Among the neoplecostomines, fin-ray strongly intumescent along its entire length with the distinctive papilla posterior to each dentary is similar to the dorsal, lateral and ventral faces ornamented with short a series of papillae found in Neoplecostomus. However, the hypertrophied odontodes, sometimes equal in size to those new species fits in the diagnosis of Pareiorhaphis proposed on head margin, that are absent in females. In addition, by Pereira et al. (2007) and can be further distinguished from fully developed males have a narrow fleshy flap along the all Neoplecostomus species by the absence of a thoracic shield entire length of the dorsal margin of the pectoral-fin spine of small dermal platelets located between the pectoral girdle E.H.L. Pereira & A.M. Zanata 41

Fig. 4. Geographic distribution and river inhabited by Pareiorhaphis lophia: (a) map of rio Paraguaçu basin, Bahia State, northeastern Brazil, showing distribution of the new species (white symbol represents type locality), (b) type locality, Lençóis, rio Paraguaçu drainage, rio Santo Antônio. and the pelvic-fin insertions, present in all Neoplecostomus also distinct from all congeners in having a small patch of species. In addition, P. lophia exhibits a single flattened distinct papillae arranged in a short median and slightly more papilla posterior to the dentaries (Fig. 2) whereas two or elevated bump (Fig. 2) at the midline of lower lip, immediately three series of large papillae are found in most species of behind the dentaries (vs. papillae not elevated in a bump). Neoplecostomus. Neoplecostomus selenae also possess one or This feature also occurs in the hypoptopomatine Hisonotus two irregular series of large flattened papillae, just posterior insperatus, which possesses a distinct median series of papillae to the dentary (Zawadzki et al., 2008), however, P. lophia at the midline of lower lip behind to the dentaries, arranged is readily distinguished from N. selenae by the presence of in a bump (Britski & Garavello, 2003; Martins & Langeani, well-developed hypertrophied odontodes and soft fleshy skin 2012). Despite of these similar morphology, H. insperatus on the lateral margin of head in adult males (vs. absence in and P. lophia are relatively distant phylogenetically and these adult males of N. selenae). Furthermore, the new species is features are likely convergent. 42 New Pareiorhaphis from northeastern Brazil

Among Pareiorhaphis species, P. bahianus described from Britski, H. A. & J. C. Garavello. 2003. Hisonotus insperatus: new species, from the upper rio Paraná basin (Pisces: Ostariophysi: Fazenda Almada, Ilhéus, is the geographically closest species Loricariidae). Copeia, 2003: 588-593. to P. lophia. However, in addition to the feature related to the Carvalho, M. & F. A. Bockmann. 2007. Família Loricariidae: papillae on lower lip and absence of dorsal-fin spinelet in most Hypostominae. Pp. 68-72. In: Buckup, P. A., N. A. Menezes & specimens, the new species further differs from P. bahianus in M. S. Ghazzi (Eds.). Catálogo das espécies de peixes de água having a smaller interorbital width and orbital diameter (25.2- doce do Brasil. Série livros 23, Museu Nacional, Rio de Janeiro. Martins, F. O. & F. Langeani. 2012. Hisonotus piracanjuba, a new 29.8 vs. 32.1-39.0% HL and 11.8-15.7 vs. 15.8-18.8% HL, species of Hypoptopomatinae (Siluriformes: Loricariidae) respectively). With the description of Pareiorhaphis lophia, from the rio Paranaíba, upper rio Paraná system, central Brazil. the genus Pareiorhaphis is now represented by two species in Ichthyological Exploration of Freshwaters, 23: 29-36. Bahia State and the new species represents the northernmost Pereira, E. H. L. 2005. Resurrection of Pareiorhaphis Miranda distribution record for the genus. Ribeiro, 1918 (Teleostei: Siluriformes: Loricariidae), and description of a new species from the rio Iguaçu basin, Brazil. Neotropical Ichthyology, 3: 271-276. Comparative material. Material examined in addition to that listed Pereira, E. H. L. & R. E. Reis. 2002. Revision of the loricariid genera by Pereira & Reis (2002), Pereira & Britto (2012) and Pereira et Hemipsilichthys and Isbrueckerichthys (Teleostei: Siluriformes), al. (2012): Pareiorhaphis proskynita. MNRJ 38561, 83.9 mm SL, with descriptions of five new species of Hemipsilichthys. holotype, Minas Gerais, rio Doce drainage, Catas Altas, ribeirão Ichthyological Exploration of Freshwaters, 13: 97-146. Pereira, E. H. L., F. Vieira & R. E. Reis. 2007. A new species of Caraça at Taboões, R. P. P. N. Santuário do Caraça, tributary to rio sexually dimorphic Pareiorhaphis Miranda Ribeiro, 1918 Piracicaba. MCP 46318, 20 paratypes, 1 c&s, 69.2 mm SL, same (Siluriformes: Loricariidae) from the rio Doce basin, Brazil. locality as the holotype. Pareiorhaphis ruschii. MCP 46640, 84.7 Neotropical Ichthyology, 5: 443-448. mm SL, holotype, Espírito Santo, Santa Teresa, córrego Lombardia Pereira, E. H. L., F. Vieira & R. E. Reis. 2010. Pareiorhaphis ca.1 km NE of the Reserva Biológica Augusto Ruschi, tributary to scutula, a new species of neoplecostomine catfish (Siluriformes: Loricariidae) from the upper rio Doce basin, Southeastern Brazil. rio Piraquê-Açu. MCP 44807, 25 paratypes, 2 c&s 53.1-61.8 mm Neotropical Ichthyology, 8: 33-38. SL, collected with the holotype. Pereira, E. H. L. & M. R. Britto. 2012. A new distinctively colored catfish of the genus Pareiorhaphis (Siluriformes: Loricariidae) Acknowledgments from the rio Doce basin, Brazil. Copeia, 2012: 519-526. Pereira, E. H. L., P. A. Lehmann & R. E. Reis. 2012. A new species of neoplecostomine catfish Pareiorhaphis (Siluriformes: We thank C. P. Oliveira, D. França, J. L. Birindelli, M. P. Loricariidae) from the coastal basins of Espírito Santo, eastern Geraldes, O. T. Oyakawa, P. Moura, P. Camelier, and R. Burger Brazil. Neotropical Ichthyology, 10: 539-546. for help in the fieldwork. The fisherman João Junior Silva da Schaefer, S. A. 1997. The neotropical cascudinhos: systematics Conceição was particularly helpful in the acquisition of adult and biogeography of the Otocinclus catfishes (Siluriformes: specimens. Priscila Camelier provided the map for Fig. 4a and Loricariidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 148: 1-120. Pablo Lehmann prepared Fig. 1. For the loan of specimens and Taylor, W. R. & G. C. Van Dyke. 1985. Revised procedures for other assistance we thank C. Lucena (MCP) and O. Oyakawa staining and clearing small fishes and other vertebrates for bone (MZUSP). Recent material was acquired in field expeditions and cartilage study. Cybium, 9: 107-119. funded by All Catfish Species Inventory (NSF DEB-0315963), Zawadzki, C. H., C. S. Pavanelli & F. Langeani. 2008. Neoplecostomus during an expedition to Paraguaçu and Itapicuru rivers (EPI), (Teleostei: Loricariidae) from the upper rio Paraná basin, Brazil, with description of three new species. Zootaxa, 1757: 31-48. in July 2005, and CNPq (# 133917/2010-0) on other occasions. Zanata, A. M., B. Sardeiro & C. H. Zawadzki. 2013. A new EHLP is partially supported by a postdoctoral fellowship dark-dotted species of Hypostomus Lacépède (Siluriformes: from CNPq (Proc. 151624/2009-7). Permission for collecting Loricariidae) from rio Paraguaçu, Bahia State, Brazil. specimens was granted by IBAMA # 13754-1. Neotropical Ichthyology, 11: 247-256.

Literature Cited Submitted August 20, 2013 Accepted December 16, 2013 by Marcelo Britto Birindelli, J. L., A. M. Zanata & F. C. T. Lima. 2007. Hypostomus Published march 31, 2014 chrysostiktos, a new species of armored catfish (Siluriformes: Loricariidae) from rio Paraguaçu, Bahia State, Brazil. Neotropical Ichthyology, 5: 271-278.